Human brain basis of musical rhythm perception: common and distinct neural substrates for meter, tempo, and pattern.

Rhythm as the time structure of music is composed of distinct temporal components such as pattern, meter, and tempo. Each feature requires different computational processes: meter involves representing repeating cycles of strong and weak beats; pattern involves representing intervals at each local time point which vary in length across segments and are linked hierarchically; and tempo requires representing frequency rates of underlying pulse structures. We explored whether distinct rhythmic elements engage different neural mechanisms by recording brain activity of adult musicians and non-musicians with positron emission tomography (PET) as they made covert same-different discriminations of (a) pairs of rhythmic, monotonic tone sequences representing changes in pattern, tempo, and meter, and (b) pairs of isochronous melodies. Common to pattern, meter, and tempo tasks were focal activities in right, or bilateral, areas of frontal, cingulate, parietal, prefrontal, temporal, and cerebellar cortices. Meter processing alone activated areas in right prefrontal and inferior frontal cortex associated with more cognitive and abstract representations. Pattern processing alone recruited right cortical areas involved in different kinds of auditory processing. Tempo processing alone engaged mechanisms subserving somatosensory and premotor information (e.g., posterior insula, postcentral gyrus). Melody produced activity different from the rhythm conditions (e.g., right anterior insula and various cerebellar areas). These exploratory findings suggest the outlines of some distinct neural components underlying the components of rhythmic structure.

Thought beyond language: neural dissociation of algebra and natural language.

A central question in cognitive science is whether natural language provides combinatorial operations that are essential to diverse domains of thought. In the study reported here, we addressed this issue by examining the role of linguistic mechanisms in forging the hierarchical structures of algebra. In a 3-T functional MRI experiment, we showed that processing of the syntax-like operations of algebra does not rely on the neural mechanisms of natural language. Our findings indicate that processing the syntax of language elicits the known substrate of linguistic competence, whereas algebraic operations recruit bilateral parietal brain regions previously implicated in the representation of magnitude. This double dissociation argues against the view that language provides the structure of thought across all cognitive domains.

Pitch discrimination in cerebellar patients: evidence for a sensory deficit.

In the last two decades, a growing body of research showing cerebellar involvement in an increasing number of nonmotor tasks and systems has prompted an expansion of speculations concerning the function of the cerebellum. Here, we tested the predictions of a hypothesis positing cerebellar involvement in sensory data acquisition. Specifically, we examined the effect of global cerebellar degeneration on primary auditory sensory function by means of a pitch discrimination task. The just noticeable difference in pitch between two tones was measured in 15 healthy controls and in 15 high functioning patients afflicted with varying degrees of global cerebellar degeneration caused by hereditary, idiopathic, paraneoplastic, or postinfectious pancerebellitis. Participants also performed an auditory detection task assessing sustained attention, a test of verbal auditory working memory, and an audiometric test. Patient pitch discrimination thresholds were on average five and a half times those of controls and were proportional to the degree of cerebellar ataxia assessed independently. Patients and controls showed normal hearing thresholds and similar performance in control tasks in sustained attention and verbal auditory working memory. These results suggest there is an effect of cerebellar degeneration on primary auditory function. The findings are consistent with other recent demonstrations of cerebellar-related sensory impairments, and with robust cerebellar auditorily evoked activity, confirmed by quantitative meta-analysis, across a range of functional neuroimaging studies dissociated from attention, motor, affective, and cognitive variables. The data are interpreted in the context of a sensory hypothesis of cerebellar function.

The boundaries of language and thought in deductive inference.

Is human thought fully embedded in language, or do some forms of thought operate independently? To directly address this issue, we focus on inference-making, a central feature of human cognition. In a 3T fMRI study we compare logical inferences relying on sentential connectives (e.g., not, or, if ... then) to linguistic inferences based on syntactic transformation of sentences involving ditransitive verbs (e.g., give, say, take). When contrasted with matched grammaticality judgments, logic inference alone recruited "core" regions of deduction [Brodmann area (BA) 10p and 8m], whereas linguistic inference alone recruited perisylvian regions of linguistic competence, among others (BA 21, 22, 37, 39, 44, and 45 and caudate). In addition, the two inferences commonly recruited a set of general "support" areas in frontoparietal cortex (BA 6, 7, 8, 40, and 47). The results indicate that logical inference is not embedded in natural language and confirm the relative modularity of linguistic processes.

Predicting judged similarity of natural categories from their neural representations.

We report a combined behavioral and functional magnetic resonance imaging (fMRI) study of conceptual similarity among members of a natural category (mammals). The study examined the relationship between computed pairwise similarity of neural responses to viewed mammals (e.g. bear, camel, dolphin) and subjective pairwise similarity ratings of the same set of mammals, obtained from subjects after the scanning session. In each functional region of interest (fROI), measures of neural similarity were compared to behavioral ratings. fROIs were identified as clusters of voxels that discriminated intact versus scrambled images of mammals (no information about similarity was used to define fROIs). Neural similarity was well correlated with behavioral ratings in fROIs covering the lateral occipital complex in both hemispheres (with overlap of the fusiform and inferior temporal gyri on the right side). The latter fROIs showed greater hemodynamic response to intact versus scrambled images of mammals whereas the fROIs that failed to predict similarity showed the reverse pattern. The findings provide novel evidence that information about the fine structure of natural categories is coarsely coded in regions of the ventral visual pathway. Implications for the theory of inductive inference are discussed.

Speech and song: the role of the cerebellum.

An exploration into cerebellar activity during the perception and production of speech and song may elucidate general underlying cerebellar functions. Recently, the cerebellum has been hypothesized to be involved with sharpening sensory input, temporal coordination and processing of motor articulation and perception, as well as instantiation of internal models that simulate the input-output characteristics of a specific system. Sung language and spoken language share many common features (physiology for articulation and perception as well as phonology, phonotactics, syntax, and semantics of the underlying language), although they differ in certain vocal and prosodic aspects. A review of the literature on perception and production of singing and speech reveals considerable overlap in the lateral aspect of the VI lobule of the posterior cerebellum, a region known to somatotopically represent the lips and tongue. This region may instantiate internal models of vocal tract articulation that simulate well learned phonological and/or segmental articulatory - auditory/orosensory mappings utilized for both speech and singing. Recent results show tendencies for left cerebellar hemispheric specialization for processing of singing and right specialization for processing of speech, both in the VI lobule of the cerebellum, inferior to that found for representing both speech and singing. Given the crossed pattern of cerebellar-cortical anatomical connectivity the findings are consistent with the hypothesis that the right cerebellum differentially processes high pass filtered information (segmental properties) and the left cerebellum differentially processes low pass filtered information (prosodic, melodic properties). Further research is necessary to examine these hypotheses and their alternatives directly.

Functional neuroanatomy of deductive inference: a language-independent distributed network.

Studies of brain areas supporting deductive reasoning show inconsistent results, possibly because of the variety of tasks and baselines used. In two event-related functional magnetic imaging studies we employed a cognitive load paradigm to isolate the neural correlates of deductive reasoning and address the role (if any) of language in deduction. Healthy participants evaluated the logical status of arguments varying in deductive complexity but matched in linguistic complexity. Arguments also varied in lexical content, involving blocks and pseudo-words in Experiment I and faces and houses in Experiment II. For each experiment, subtraction of simple from complex arguments (collapsing across contents) revealed a network of activations disjoint from regions traditionally associated with linguistic processing and also disjoint from regions recruited by mere reading. We speculate that this network is divided into "core" and "support" regions. The latter include left frontal (BA 6, 47) and parietal (BA 7, 40) cortices, which maintain the formal structure of arguments. Core regions, in the left rostral (BA 10p) and bilateral medial (BA 8) prefrontal cortex, perform deductive operations. Finally, restricting the complex-simple subtraction to each lexical content uncovered additional activations which may reflect the binding of logical variables to lexical items.

Music and language side by side in the brain: a PET study of the generation of melodies and sentences.

Parallel generational tasks for music and language were compared using positron emission tomography. Amateur musicians vocally improvised melodic or linguistic phrases in response to unfamiliar, auditorily presented melodies or phrases. Core areas for generating melodic phrases appeared to be in left Brodmann area (BA) 45, right BA 44, bilateral temporal planum polare, lateral BA 6, and pre-SMA. Core areas for generating sentences seemed to be in bilateral posterior superior and middle temporal cortex (BA 22, 21), left BA 39, bilateral superior frontal (BA 8, 9), left inferior frontal (BA 44, 45), anterior cingulate, and pre-SMA. Direct comparisons of the two tasks revealed activations in nearly identical functional brain areas, including the primary motor cortex, supplementary motor area, Broca's area, anterior insula, primary and secondary auditory cortices, temporal pole, basal ganglia, ventral thalamus, and posterior cerebellum. Most of the differences between melodic and sentential generation were seen in lateralization tendencies, with the language task favouring the left hemisphere. However, many of the activations for each modality were bilateral, and so there was significant overlap. While clarification of this overlapping activity awaits higher-resolution measurements and interventional assessments, plausible accounts for it include component sharing, interleaved representations, and adaptive coding. With these and related findings, we outline a comparative model of shared, parallel, and distinctive features of the neural systems supporting music and language. The model assumes that music and language show parallel combinatoric generativity for complex sound structures (phonology) but distinctly different informational content (semantics).

The neural basis of human dance.

Human dance was investigated with positron emission tomography to identify its systems-level organization. Three core aspects of dance were examined: entrainment, meter and patterned movement. Amateur dancers performed small-scale, cyclically repeated tango steps on an inclined surface to the beat of tango music, without visual guidance. Entrainment of dance steps to music, compared to self-pacing of movement, was supported by anterior cerebellar vermis. Movement to a regular, metric rhythm, compared to movement to an irregular rhythm, implicated the right putamen in the voluntary control of metric motion. Spatial navigation of leg movement during dance, when controlling for muscle contraction, activated the medial superior parietal lobule, reflecting proprioceptive and somatosensory contributions to spatial cognition in dance. Finally, additional cortical, subcortical and cerebellar regions were active at the systems level. Consistent with recent work on simpler, rhythmic, motor-sensory behaviors, these data reveal the interacting network of brain areas active during spatially patterned, bipedal, rhythmic movements that are integrated in dance.

The brain basis of piano performance.

Performances of memorized piano compositions unfold via dynamic integrations of motor, perceptual, cognitive, and emotive operations. The functional neuroanatomy of such elaborately skilled achievements was characterized in the present study by using (15)0-water positron emission tomography to image blindfolded pianists performing a concerto by J.S. Bach. The resulting brain activity was referenced to that for bimanual performance of memorized major scales. Scales and concerto performances both activated primary motor cortex, corresponding somatosensory areas, inferior parietal cortex, supplementary motor area, motor cingulate, bilateral superior and middle temporal cortex, right thalamus, anterior and posterior cerebellum. Regions specifically supporting the concerto performance included superior and middle temporal cortex, planum polare, thalamus, basal ganglia, posterior cerebellum, dorsolateral premotor cortex, right insula, right supplementary motor area, lingual gyrus, and posterior cingulate. Areas specifically implicated in generating and playing scales were posterior cingulate, middle temporal, right middle frontal, and right precuneus cortices, with lesser increases in right hemispheric superior temporal, temporoparietal, fusiform, precuneus, and prefrontal cortices, along with left inferior frontal gyrus. Finally, much greater deactivations were present for playing the concerto than scales. This seems to reflect a deeper attentional focus in which tonically active orienting and evaluative processes, among others, are suspended. This inference is supported by observed deactivations in posterior cingulate, parahippocampus, precuneus, prefrontal, middle temporal, and posterior cerebellar cortices. For each of the foregoing analyses, a distributed set of interacting localized functions is outlined for future test.

The role of the insular cortex in pitch pattern perception: the effect of linguistic contexts.

Auditory pitch patterns are significant ecological features to which nervous systems have exquisitely adapted. Pitch patterns are found embedded in many contexts, enabling different information-processing goals. Do the psychological functions of pitch patterns determine the neural mechanisms supporting their perception, or do all pitch patterns, regardless of function, engage the same mechanisms? This issue is pursued in the present study by using 150-water positron emission tomography to study brain activations when two subject groups discriminate pitch patterns in their respective native languages, one of which is a tonal language and the other of which is not. In a tonal language, pitch patterns signal lexical meaning. Native Mandarin-speaking and English-speaking listeners discriminated pitch patterns embedded in Mandarin and English words and also passively listened to the same stimuli. When Mandarin listeners discriminated pitch embedded in Mandarin lexical tones, the left anterior insular cortex was the most active. When they discriminated pitch patterns embedded in English words, the homologous area in the right hemisphere activated as it did in English-speaking listeners discriminating pitch patterns embedded in either Mandarin or English words. These results support the view that neural responses to physical acoustic stimuli depend on the function of those stimuli and implicate anterior insular cortex in auditory processing, with the left insular cortex especially responsive to linguistic stimuli.

Passive music listening spontaneously engages limbic and paralimbic systems.

In this PET study, non-musicians passively listened to unfamiliar instrumental music revealed afterward to elicit strongly pleasant feelings. Activations were observed in the subcallosal cingulate gyrus, prefrontal anterior cingulate, retrosplenial cortex, hippocampus, anterior insula, and nucleus accumbens. This is the first observation of spontaneous responses in such limbic and paralimbic areas during passive listening to unfamiliar although liked music. Activations were also seen in primary auditory, secondary auditory, and temporal polar areas known to respond to music. Our findings complement neuroimaging studies of aesthetic responses to music that have used stimuli selected by subjects or designed by experimenters. The observed pattern of activity is discussed in terms of a model synthesizing emotional and cognitive responses to music.

The song system of the human brain.

Although sophisticated insights have been gained into the neurobiology of singing in songbirds, little comparable knowledge exists for humans, the most complex singers in nature. Human song complexity is evidenced by the capacity to generate both richly structured melodies and coordinated multi-part harmonizations. The present study aimed to elucidate this multi-faceted vocal system by using 15O-water positron emission tomography to scan "listen and respond" performances of amateur musicians either singing repetitions of novel melodies, singing harmonizations with novel melodies, or vocalizing monotonically. Overall, major blood flow increases were seen in the primary and secondary auditory cortices, primary motor cortex, frontal operculum, supplementary motor area, insula, posterior cerebellum, and basal ganglia. Melody repetition and harmonization produced highly similar patterns of activation. However, whereas all three tasks activated secondary auditory cortex (posterior Brodmann Area 22), only melody repetition and harmonization activated the planum polare (BA 38). This result implies that BA 38 is responsible for an even higher level of musical processing than BA 22. Finally, all three of these "listen and respond" tasks activated the frontal operculum (Broca's area), a region involved in cognitive/motor sequence production and imitation, thereby implicating it in musical imitation and vocal learning.

Left and right hand recognition in upper limb amputees.

Previous research suggests a close similarity in brain activity between mental simulation of a movement and its real counterpart. To explore this similarity, we aimed to assess whether imagery is affected by the loss of a limb or of its motor skills. We examined the performance of 16 adult, upper limb amputees (and age-matched controls) in a left/right hand judgement task that implicitly requires motor imagery. The experimental group included subjects who had suffered the amputation of the dominant or the non-dominant limb. Although responding well above chance, amputees as a group were slower and less accurate than controls. Nevertheless, their response pattern was similar to that of controls, namely slower response times and more errors for stimuli depicting hands in unnatural orientations, i.e. postures difficult to reach with a real movement. Interestingly, for all stimuli, amputees' performance was strongly affected by the side of limb loss: subjects who underwent amputation of their preferred limb made more errors and required greater latencies to respond as compared with amputees of the non-dominant limb. In a further analysis we observed that the habit of wearing an aesthetic prosthesis significantly interfered with the ability to judge the corresponding hand. Our data lead to three main conclusions: (i) loss of a single limb per se does not prevent motor imagery but it significantly enhances its difficulty; (ii) these subjects apparently perform the hand recognition task using a strategy in which they initially mentally simulate movements of their dominant limb; (iii) wearing a prosthesis, devoid of any motor function, seems to interfere with motor imagery, consistent with the view that only 'tools' can be incorporated in a dynamic body schema.

Superior parietal cortices and varieties of mental rotation.

Mental rotation is the most distinctly specialized operation of the imagination, one characterized precisely enough psychophysically for parametric study, thereby making it an optimal prospect for isolating and modeling its neural mechanisms. New human brain mapping studies using direct cortical stimulation and repetitive transcranial magnetic stimulation isolate an area in right superior parietal cortex that appears to be crucial for the mental rotation of objects, but not to the mental rotation of one's body.